TRAWL CAUGHT IN THE SOUTHERN BIGHT : a guide to field and laboratory identification

Donald B. Cadien, CSDLAC, 3 April 1998

Because they are prominent in many trawl catches several comprehensive regional treatments of shrimp exist. The most pertinent are Schmitt (1921), Word and Charwat (1976) and Butler (1980). Each of these had a slightly different focus, and none is adequate for all the shrimp recorded from the Bight. Martin & Zimmerman (1997) cover the northern portion of the Santa Barbara Channel, but treat only eleven species. Butler provides the most extended descriptions of the species covered, but treats the fauna of the Pacific Northwest and lacks many southern species. Word and Charwat focussed on the Bight, using the same sort of monitoring records we currently do. Nomenclatural changes have reduced the utility of their treatment, and although nearly all species are keyed, little supporting description is supplied. Schmitt likewise offers little detail in his descriptions of the species. Taxonomic changes have rendered many of his names obsolete as well. SCAMIT Taxonomic List Ed. 3 will provide the synonymies which allow older usages like Schmitt’s to be related to current usage. Other resources are available for particular groups, especially the series of papers by Wicksten dealing with southern California families, genera, or species (Wicksten 1976, 1977, 1978a, b; 1979; 1980; 1981; 1983a; b; 1984; 1986; 1989a, b; 1990a, b; 1991, 1992, 1996a, b; Wicksten & Butler 1983; Wicksten & Hendrickx 1991). Live appearance of many shrimp is documented by color photographs in Jensen 1995. Butler 1980 gives color drawings of many species from depths inaccessible to diving photographers. Species which are normally found to the south, but have made incursions into the Southern California Bight during the current strong ENSO event require additional references (Chace 1937; Hendrickx 1990, 1995, 1996; Hendrickx & Navarrete 1996; Hendrickx & Wicksten 1989).

Because such a variety of information sources is available on this group, the necessity of precalibration of trawl shrimp identification in the Bight ‘98 regional sampling effort is great. With that goal in mind, new keys to known from the Bight which can be identified in the field have been prepared where necessary. Members of the and the do not easily lend themselves to field separation because of size and/or use of small or obscure character states in their identification. Members of these families should always be returned to the lab for identification. Other groups such as the may or may not be field separable depending on the experience and expertise of the observer. In such intermediate groups it is particularly important to recognize your own limitations. A good rule of thumb for deciding if specimens should be returned to the lab for further identification is "if you have ANY doubts as to the identity of the , they should be returned to the lab for confirmation." Not just a few representatives, BUT ALL SUSPECT SPECIMENS. Even in cases where the observer has no doubt as to the identity of the animals, vouchers must still be taken for lab confirmation. These should be collected by each individual who participates in the field identifications for each species they have identified.

These requirements are not new, but they must be followed by all participants to avoid data compromise. If only one participant fails to identify, or identifies incorrectly, material which has been discarded prior to the discovery of the data deficiency, all data provided by other participants must be degraded to the level of the non-conforming group prior to analysis. This unfortunately occurred on several occasions during the 1994 regional monitoring effort, and the utility of resulting data was diminished. These concerns become even greater during Bight ‘98 because the number of groups involved is being increased, and along with the number of participants the possibilities for non-conformity.

The following families of shrimp are known from historic records to have occurred in the Bight - Alpheidae, Aristaeidae, Crangonidae, , Hippolytidae, Luciferidae, Ogyrididae, , , , , Penaeidae, , Sergestidae, Sicyoniidae, and Solenoceridae (less than half the 36 known families of shrimps). Of these sixteen families the Aristaeidae, Luciferidae, Oplophoridae, Pasiphaeidae and Sergestidae are all holopelagic shrimps and not part of the bottom trawl fauna.

For those interested in these animals a list of species reported from our area and useful references for their identification are provided below.

Aristaeidae Bentheogennema burkenroadi Krygier & Wasmer 1975 has been taken from the surface to 1000m depths Bentheogennema borealis (Rathbun 1902) seldom reaches depths as shallow as 200m. These two species are keyed, described, and illustrated in Butler 1980. Should you reach Aristaeidae in the Key to Shrimp Families you should consult Butler for further information.

Luciferidae typus H. Milne Edwards 1837 known from as far north as the middle of the Baja peninsula, this species may range into Bight waters on the northward ENSO flow. See Hendrickx & Navarrete 1996

Oplophoridae - Species of , Systellaspis, and are known to range into our geographic coverage area, but all from deeper than our maximum depths. It is very unlikely that they will stray into our depth range. Although not yet reported from Californian waters, the Notostomus has been recorded as far south as Oregon (Butler 1980). Animals keying to Oplophoridae should be examined with Butler in hand. He provides both generic and specific keys to local species, as well as good illustrations and descriptions.

Pasiphaeidae - most species listed in Word & Charwat and Hendrickx & Navarrete fall outside the geographic or bathymetric limits of our study area. Pasiphaea pacifica Rathbun 1902 known throughout the bight, but generally taken in deeper trawls, although may surface at night (0-1076m). See Butler 1980 Pasiphaea chacei Yaldwyn 1962 from off to off Oregon, generally deeper than 300 m. Given the dislocations caused by the ENSO event, these are probably all below our depth range at this time. We should be aware of the characters to check to distinguish this from P. pacifica. See keys in Word & Charwat 1976 and Hendrickx & Navarrete 1996.

Sergestidae - Species in the genus Sergia, while taken from the surface to over 1000m depth, are from oceanic water masses found outside the Bight in the Eastern Pacific, and are not considered here. The single reported species of the genus Petalidium to occur off southern California is similarly oceanic, and not recorded from inshore waters of the Bight. Both genera are covered by Hendrickx & Navarrete 1996, who provide a key to the genera as well.

Sergestes similis Hanson 1903 ranges from the Gulf of California to the Gulf of Alaska and from the surface to 1200 m - the only member of the genus to occur in inshore waters of the Bight. Other species may occur further offshore in the California Current or beyond (see Hendrickx & Navarrete 1996). The species is well described and illustrated by Butler 1980.

Of the remaining 11 families known from the area three are penaeoids (Solenoceridae, Sicyoniidae, and Penaeidae) all at one time considered to be subfamilies within the Penaeidae. If additional information on penaeoid biology or morphology is desired consult Dall et al (1990). A separate key is provided for benthic members of these families known to occur within the Bight plus a very similar species not yet known from the area (see attached key).

The eight benthic families of caridoid shrimp known from Southern California waters are all included in the family key. All members of the Glyphocrangonidae occur too deep to fall within our coverage. Distributional records are provided by Wicksten (1979), and a good illustrated key by Hendrickx (1995). All local members of the family Ogyrididae occur too shallowly to fall within our coverage. Although Ogyrides alphaerostris was reported from the area by Wicksten & Hendrickx (1991), the local species is still undescribed. Information on it should be sought from Jim Roney (LACEMD), who is in the process of describing these shrimp. We will probably see these small burrowing shrimp in our shallowest benthic samples, but not in our trawls.

Two of the six remaining families (Processidae and Palaemonidae) are represented by only a few species. The Processidae have only two local representatives, Ambidexter panamensis and Processa peruviana. Ambidexter has been reported only from shallow water in San Diego Bay, where it is taken both in infaunal samples and in seines. It is apparently a burrowing species, and may only be taken during night or crepuscular samplings over mud/algal bottoms. Given the dearth of records, the population in the bay may be quite localized. Abele (1972) describes the . If shallow water samples are taken in San Diego Bay we might get this species. Processa peruviana has only been taken once in local waters; off Palos Verdes in 1995. This animal is larger than A. panamensis, and was taken in the open sea, not in a bay, where it favors fine sand bottoms (Hendrickx 1995). It was taken at night, and may also be a burrower. Wicksten’s original description (1983), supplemented by information in Hendrickx (1995), should allow identification of further specimens. Hendrickx also provides a key to separate these two genera. The normal range of the animal extends only as far north as the tip of Baja California, so occurrence in our area is undoubtedly related to ENSO transport.

Local Palaemonidae can be adequately field separated with the key of Word & Charwat (1976), which covers all species recorded to date. If you reach Palaemonidae in the family key, consult the above key. Members of the genus Palaemon occur only in shallow estuarine areas such as Huntington Harbor. Although an indigenous species exists (Palaemon ritteri), all recently caught Palaemon have been the introduced P. macrodactylus, which would key to the same place in Word & Charwat’s key. Of the remaining four species in the key, two are commensals and not likely to be taken in a trawl sample. Pontonia californiensis is an endocommensal of tunicates, living within the branchial basket of the host. Pseudocoutierea elegans is an obligate commensal of muricid sea-fans, and may abandon a host caught in a trawl net and swim away. Specimens are usually taken from sea-fans collected by divers, but could also come from trawls which take sea-fans. The remaining species, Palaemonella holmesi and Periclimenes infraspinus, though rarely taken, could be caught in shallow water trawls within the Bight. Palaemonetes hiltoni, described from San Pedro by Schmitt in 1921 has not been seen since, and is no longer considered to occur in the Bight (Wicksten 1989). Under the current ENSO transport regime, we may refind it or other southern palaemonid species which have not yet been reported here. Wicksten’s key (1989) to the family includes species not yet known from the Bight, and should be consulted for specimens which appear to key poorly or not at all in Word & Charwat 1976. Any specimens keying to the family should probably be retained for laboratory verification. The two-volume monograph by Holthuis (1951, 1952) should provide information on species (such as Palaemon macrodactylus) not described in other works.

Members of the families Alpheidae and Hippolytidae should always be returned to the laboratory for identification, or for verification if large and characteristic enough for field ID. No specimens of these families should be discarded in the field. The sole exception to this rule is the hippolytid californica. This species is large enough, and has a characteristic enough live appearance (see photograph in Jensen 1995) for reliable determination in the field. Voucher specimens should still be returned to the lab for verification, but large collections of L. californica can be returned to the sea. Although nearly all hippolytid species can be correctly determined with the key in Word and Charwat, we should use that in Wicksten (1990). California alpheids can be separated using the attached key, which includes undescribed species not in Wicksten (1984).

Species in the families Crangonidae and Pandalidae should be identifiable in the field in nearly all cases. Some damaged specimens and some very small juveniles may require laboratory confirmation, but few specimens should fall into these categories. New keys to both these families are presented here to incorporate taxonomic changes and new species records which render existing keys incomplete.

Resource Guide - Recommended Standard References for Bight ‘98 Trawl Shrimp Identification F= field key, L= laboratory key, C= combined field and laboratory key

Trawl Shrimp Families - attached Family key (C) - all families below are included in the key

Alpheidae - LABORATORY ID ONLY, collect all specimens - attached Alpheid key (L) Aristaeidae - all members excluded as holopelagic Crangonidae - attached Crangonid key (C) Glyphocrangonidae - all members excluded as outside depth limits Hippolytidae - LABORATORY ID ONLY, collect all specimens - Wicksten 1990b key (L) Luciferidae - all members excluded as holopelagic Ogyrididae - all members excluded as outside depth limits Oplophoridae - all members excluded as holopelagic Palaemonidae - Word & Charwat 1976 key (F), Wicksten 1989a key (L) Pandalidae - attached Pandalid key (C) Pasiphaeidae - all members excluded as holopelagic Penaeidae - attached Penaeoid key (C), Hendricks 1995 key (L) Processidae - Hendrickx 1995 key (C) Sergestidae - all members excluded as holopelagic Sicyoniidae - attached Penaeoid key (C) Solenoceridae - attached Penaeoid key (C) List of Species Taken in Association with Monitoring Programs in the Southern California Bight at depths between 10-300 m

Suborder Penaeidea Family Hippolytidae Superfamily Eualus herdmani (Walker 1898) Family Aristeidae Eualus lineatus Wicksten & Butler 1983 Bentheogennema burkenroadi Krygier & Wasmer 1975 Heptacarpus brevirostris (Dana 1852) Family Solenoceridae Heptacarpus decorus (Rathbun 1902) mutator Burkenroad 1938 Heptacarpus fuscimaculatus Wicksten 1986 Family Penaeidae Heptacarpus flexus (Rathbun 1902) Metapenaeopsis mineri Burkenroad 1934 Heptacarpus palpator (Owen 1839) Penaeus californiensis Holmes 1900 Heptacarpus sitchensis (Brandt 1851) Family Sicyoniidae Heptacarpus stimpsoni Holthuis 1947 ingentis (Burkenroad 1938) Heptacarpus taylori (Stimpson 1857) Sicyonia penicillata Lockington 1879 Heptacarpus tenuissimus Holmes 1900 Superfamily Heptacarpus tridens (Rathbun 1902) Family Sergestidae Hippolyte californiensis Holmes 1895 Sergestes similis Hansen 1903 Hippolyte clarki Chace 1951 Suborder Lysmata californica (Stimpson 1866) Superfamily Pasiphaeoidea Spirontocaris holmesi Holthuis 1947 Family Pasiphaeidae Spirontocaris lamellicornis (Dana 1852) Pasiphaea pacifica Rathbun 1902 Spirontocaris prionota (Stimpson 1864) Superfamily Spirontocaris sica Rathbun 1902 Family Pandalidae Spirontocaris snyderi Rathbun 1902 Pandalopsis ampla Bate 1888 Family Ogyrididae danae Stimpson 1857 Ogyrides sp A Roney 1978 § Pandalus jordani Rathbun 1902 Family Processidae Brandt 1851 Processa peruviana Wicksten 1983 Pantomus affinis Chace 1937 Superfamily Plesionika beebei Chace 1937 Family Crangonidae Plesionika trispinus Squires & Barragán 1976 Crangon alaskensis Lockington 1877 Superfamily Crangon alba Holmes 1900 Family Alpheidae Crangon handi Kuris & Carlton 1977 Alpheopsis equidactylus (Lockington 1877) Crangon holmesi Rathbun 1902 Alpheus bellimanus Lockington 1877 Crangon nigricauda Stimpson 1856 Alpheus californiensis Holmes 1900 Crangon nigromaculata Lockington 1877 Alpheus clamator Lockington 1877 Mesocrangon munitella (Walker 1898) Automate sp A SCAMIT 1995 § Metacrangon spinosissima (Rathbun 1902) Betaeus ensenadensis Glassell 1938 Neocrangon communis (Rathbun 1899) Betaeus harfordi (Kingsley 1878) Neocrangon resima (Rathbun 1902) Betaeus harrimani Rathbun 1904 Neocrangon zacae (Chace 1937) Betaeus longidactylus Lockington 1877 Rhynocrangon alata (Rathbun 1902) Superfamily Family Palaemonidae Pseudocoutierea elegans Holthuis 1951

BIBLIOGRAPHY

ABELE, LAWRENCE G. 1972. A review of the genus Ambidexter (Crustacea: : Processidae) in Panama. Bulletin of Marine Science 22(2):365-380. BUTLER, TERRANCE H. 1980. Shrimps of the Pacific coast of Canada. Canadian Bulletin of Fisheries and Aquatic Sciences (202):1-280. CHACE, FENNER A. Jr. 1937. The Templeton Crocker Expedition. VII. Caridean decapod Crustacea from the Gulf of California and the West Coast of Lower California. Zoologica 22(2):109-138. DALL, W., B. J. Hill, P. C. Rothlisberg, D. J. Staples. 1990. Advances in Marine Biology Volume 27. The Biology of the Penaeidae. Academic Press Limited: London. 489pp. HENDRICKX, MICHEL E. 1990. The stomatopod and decapod collected during the GUAYTEC II Cruise in the Central Gulf of California, Mexico, with the description of a new species of Plesionika Bate (Caridea: Pandalidae). Revista de Biologia Tropical 38(1): 35-53. ---. 1995. Camarones. Pp.417-537. IN: Fischer, W., F. Krupp, W. Schneider, C. Sommer, Carpenter K. E., and V. H. Niem (eds.), Guia FAO para la identificación de especies para los fines de la pesca. Pacifico centro-Oriental. Vol. 1. Plantas e Invertebrados. F.A.O.: Rome, Italy. 646pp. ---. 1996. Los Camarones Penaeoidea Bentónicos (Crustacea: Decapoda: ) del Pacífico Mexicano. 147pp. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad e Instituto de Ciencias del Mar y Limnologia, UNAM: Guadalajara, Mexico. HENDRICKX, MICHEL E., and Flor D. Estrada-Navarrete. 1989. A checklist of the species of pelagic shrimps (Penaeoidea and Caridea) from the Eastern Pacific, with notes on their geographic and depth distribution. CalCOFI Report 30:104-121. ---. 1996. Los Camarones Pelágicos (Crustacea: Dendrobranchiata y Caridea) del Pacífico Mexicano. 157pp. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad e Instituto de Ciencias del Mar y Limnologia, UNAM: Guadalajara, Mexico. HENDRICKX, MICHEL E., and Mary K. Wicksten. 1989. Los Pandalidae (Crustacea: Caridea) del Pacifico Mexicano, con una clave para su identificacion. Caldasia 16(76):71-86. HOLTHUIS, LIPKE B. 1951. A general revision of the Palaemonidae (Crustacea Decapoda Natantia) of the Americas. I. The subfamilies Euryrhynchinae and Pontoniinae. Allan Hancock Foundation Publications, Occasional Paper (11):1-332. ---. 1952. A general revision of the Palaemonidae (Crustacea Decapoda Natantia) of the Americas. II. Subfamily Palaemoninae. Allan Hancock Foundation Publications, Occasional Paper (12):1- 396. JENSEN, GREGORY C. 1995. Pacific Coast Crabs and Shrimps. Sea Challengers: Monterey, California. 87pp. MARTIN, JOEL W., and Todd L. Zimmerman. 1997. Order Decapoda. Pp. 49-121 IN: Blake, James A. And Paul H. Scott (eds.). Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 10 The Arthropoda - The Pycnogonida, The Crustacea Part 1 - The Decapoda and Mysidacea. Santa Barbara Museum of Natural History, Santa Barbara, California. 151pp. SCHMITT, WALDO L. 1921. The Marine Decapod Crustacea of California, with special reference to the Decapod Crustacea collected by the United States Bureau of Fisheries Steamer 'Albatross' in connection with the Biological Survey of San Francisco Bay during the years 1912-1913. University of California Publications in Zoology 23:1-470. WICKSTEN, MARY K. 1976. First record of Argis levior (Rathbun) from California (Decapoda: Crangonidae). Bulletin of the Southern California Academy of Sciences 75(1):56. ---. 1977. Range extentions of four species of crangonid shrimps from California and Baja California, with a key to the genera (Natantia: Crangonidae). Proceedings of the Biological Society of Washington 90( ):963-967 ---. 1978a. The species of Plesionika from California and Western Mexico (Natantia: Pandalidae). Bulletin of the Southern California Academy of Sciences 77(2):84-87. ---. 1978b. The species of Lebbeus in California (Crustacea: Caridea: Hippolytidae). Occasional Papers of the Allan Hancock Foundation, new series (1):1-8. ---. 1979. New records of the species of Glyphocrangon in the northeastern Pacific Ocean (Caridea: Glyphocrangonidae). Proceedings of the Biological Society of Washington 92(2):217- 224. ---. 1980. Range extensions of four species of crangonid shrimps in the eastern Pacific Ocean (Decapoda: Crangonidae). Bulletin of the Southern California Academy of Sciences 79(1):38-41. ---. 1981. The species of Automate (Caridea: Alpheidae) in the eastern Pacific Ocean. Proceedings of the Biological Society of Washington 94(4):1102-1109. ---. 1983a. A monograph on the shallow water caridean shrimps of the Gulf of California, Mexico. Allan Hancock Monographs in Marine Biology (13):1-59. ---. 1983b. Plesionika sanctaecatalinae: a new species of deep- sea shrimp from the eastern Pacific (Caridea: Pandalidae). Bulletin of the Southern California Academy of Sciences 82(3): 138-143. ---. 1984. New records of snapping shrimps (Family Alpheidae) from California. Proceedings of the Biological Society of Washington 97(1):186-190. ---. 1986. A new species of Heptacarpus from California, with a redescription of Heptacarpus palpator (Owen) (Caridea: Hippolytidae). Bulletin of the Southern California Academy of Sciences 85(1):46-55. ---. 1989a. A key to the palaemonid shrimp of the Eastern Pacific Region. Bulletin of the Southern California Academy of Sciences 88(1):11-20. ---. 1989b. Ranges of offshore decapod crustaceans in the eastern Pacific Ocean. Transactions of the San Diego Society of Natural History 21(19):291-316. ---. 1990a. On the status of Alpheus barbara Lockington (Crustacea: Caridea: Alpheidae). Proceedings of the Biological Society of Washington 103(1):100-102. ---. 1990b. Key to the hippolytid shrimp of the Eastern Pacific Ocean. Fishery Bulletin 88(3):587-598. ---. 1991. Pandalus gurneyi Stimpson synonymized with Pandalus danae Stimpson (Decapoda: Pandalidae). Proceedings of the Biological Society of Washington 104(4):812-815. ---. 1992. A new species of Alpheopsis (Decapoda, Alpheidae) from the tropical eastern Pacific, with a key to the species of Alpheopsis of the Americas. Crustaceana 63(1):51-56. ---. 1996a. Heptacarpus pictus (Stimpson) synonymized with Heptacarpus sitchensis (Brandt)(Decapoda, Hippolytidae). Crustaceana 69(1):71-75. ---. 1996b. Neocrangon zacae (Chace, 1937) synonymized with N. resima (Rathbun, 1902), and compared with N. communis (Rathbun, 1899) (Decapoda: Caridea: Crangonidae). Proceedings of the Biological Society of Washington 109(1):39-43. WICKSTEN, MARY K., and Terrance H. Butler. 1983. Description of Eualus lineatus new species, with a redescription of Heptacarpus herdmani (Walker) (Caridea: Hippolytidae). Proceedings of the Biological Society of Washington 96(1):1-6. WICKSTEN, MARY K., and Michel E. Hendrickx. 1991. Checklist of penaeoid and caridean shrimps (Decapoda: Penaeoidea, Caridea) from the eastern tropical Pacific. Proceedings of the San Diego Society of Natural History 9:1-11. WORD, JACK Q., and Danuta Charwat. 1974. Key to shrimp common in Southern california trawl catches. Southern California Coastal Water Research Project (SCCWRP), Technical Memorandum (221):1-41. KEY TO SHRIMP FAMILIES RECORDED IN THE SOUTHERN CALIFORNIA BIGHT D. B. Cadien (CSDLAC) - 7Dec1994 (rev. 25Mar1998) (based on Chace 1972, Burukovskii 1974, Dall et al 1990, and Hendrickx 1995)

1. Third legs chelate; pleura of second abdominal somite not overlapping that of first somite; ...... Suborder Dendrobranchiata 2 Third legs not chelate; pleura of second abdominal somite overlapping that of first somite; ...... Suborder Eukyphida 7 2. Legs 4 and 5 well developed; gills numerous ...... Superfamily Penaeoidea 4 Legs 4 and 5 reduced or absent; gills few (<9) ...... Superfamily Sergestoidea 3 3. Legs 4 and 5 absent; carapace elongated anteriorly by narrow "neck", eyes widely separated from mouthparts ...... Luciferidae Legs 4 and 5 reduced; carapace not elongated anteriorly, eyes just slightly anterior to mouthparts ...... Segestidae 4. Antennular flagellae subequal and originate distally on the 3rd segment; 5th leg lacks epipod...... 5 Median(upper) flagellum much shorter than lateral (lower) and originates near base of 3rd segment; 5th leg with an epipod...... Aristaeidae 5 Cervical sulcus reaching < the distance from the hepatic spine to the top of the carapace; postorbital spine absent;4th leg lacks epipod...... 6 Cervical sulcus reaching the top of the carapace; postorbital spine present; 4th leg with an epipod ...... Solenoceridae 6. Third to 5th pleopods each with 2 rami; prosartema (eye brush) present on antennula; exopods present on 2nd and 3rd maxillipeds ...... Penaeidae Third to 5th pleopods each with a single ramus; prosartema absent; no exopods on 2nd and 3rd maxillipeds ...... Sicyoniidae 7. First leg subchelate...... 8 First leg chelate or simple ...... 9 8. Carpus of 2nd leg unsegmented...... Crangonidae Carpus of 2nd leg multisegmented...... Glyphocrangonidae 9. First and 2nd legs chelate; fingers of chelae with pectinate edges ...... Pasiphaeidae First and/or 2nd legs chelate; fingers without pectinate edges...... 10 10.Carpus of 2nd leg unsegmented; 1st leg with well-developed chela...... 11 Carpus of 2nd leg multisegmented, OR 1st leg not chelate...... 12 11.Legs with exopods...... Oplophoridae Legs lacking exopods...... Palaemonidae 12.First legs with at least one well developed chela...... 13 First legs with chelae very small or absent...... Pandalidae 13.Rostrum edentate or dentate, but without subdistal tooth ...... 14 Rostrum with distal notch covered with bristles and forming subdistal dorsal tooth...... Processidae 14. Eyes on long stalks, reaching nearly to end of antennular peduncle, and several times longer than eye diameter ...... Ogyrididae Eyestalks not unusually long, not or only slightly exceeding eye diameter ...... 15 15. Eyes usually partially or entirely covered by carapace, incapable of free lateral movement; rostrum absent or spinelike...... Alpheidae Eyes exposed and freely movable; rostrum well developed, toothed...... Hippolytidae KEY TO THE SO. CALIFORNIA BIGHT CRANGONID SHRIMP Donald B. Cadien (CSDLAC), 20 March 1998 (based on Word & Charwat 1976, Kuris & Carlton 1977, Wicksten 1977, and Butler 1980)

1. Dactyls of 4th and 5th legs flattened; eyes partly concealed by carapace ...... Argis 2 Dactyls of 4th and 5th legs not flattened; eyes not partly concealed...... 3

2. Carapace with 2 median dorsal spines posterior to rostral spine...... Argis californiensis Carapace with 4 median dorsal spines posterior to rostral spine...... Argis levior

3. Abdominal somites heavily sculptured; 3rd abdominal somite with rostrate posterior margin, 4th and 5th with posteriomedial spine...... Rhynocrangon alata Abdominal somites weakly sculptured or unsculptured; 3rd abdominal somite not posteriorly produced, 4th and 5th somites lacking posteromedial spine (may be carinate)...... 4

4. Gastric region of carapace depressed...... 5 Gastric region not depressed below general level of carapace...... 7

5. 1-2 spines ventrally on abdominal pleura...... Metacrangon spinosissima Abdominal pleura without ventral spines...... 6

6. Carapace lacking lower submedian spine...... Metacrangon munita Carapace with lower submedian spine...... Mesocrangon munitella

7. Carapace with 1 median dorsal spine ...... Crangon 8 Carapace with 2 median dorsal spines...... Neocrangon 13

8. Sixth abdominal somite with bold lateral blue pigment spot(s)...... Crangon nigromaculata Sixth abdominal somite lacking lateral pigment spot...... 9

9. Inner flagellum of antenna one distinctly longer than outer ...... 10 Inner and outer flagella of antenna one of equal length...... Crangon handi

10. Sixth abdominal somite grooved ventrally ...... 11 Sixth abdominal somite not grooved ventrally ...... 12

11. Spine of antennal scale extending well past end of blade...... Crangon alaskensis Spine of antennal scale not or barely extending past end of blade...... Crangon nigricauda

12. Ischium of 3rd maxilliped flattened and laterally flanged ...... Crangon alba Ischium of 3rd maxilliped not especially flattened and not flanged ...... Crangon holmesi

13. Third through fifth abdominal somites dorsally carinate...... Neocrangon communis Abdominal somites not dorsally carinate...... 14

14. Rostrum bearing terminal "moustache" of setae projecting obliquely downward; rostrum variable, typically horizontal or slightly elevated ...... Neocrangon zacae Rostrum with lateral field of setae, but lacks "moustache"; rostrum variable, typically bearing a ventral blade, and strongly elevated above horizontal ...... Neocrangon resima REVISED KEY TO THE PANDALIDAE OF CALIFORNIA Donald B. Cadien (CSDLAC) - 20 Mar 1998 (modified from Burukovskii 1974, Wicksten 1978, Butler 1980, and Hendrickx 1996)

1. Rostrum articulated to front of carapace...... Pantomus affinis Rostrum not articulated, integral to carapace ...... 2

2. Third maxilliped with an exopod ...... Plesionika 3 Third maxilliped lacking exopod ...... 6

3. Second legs markedly unequal in length ...... Plesionika mexicana Second legs equal or subequal in length ...... 4

4. Rostrum with 2-8 small spines dorsally near it’s base; ventral spines (if present) very small and restricted to the distal half of the rostrum...... Plesionika sanctacatalinae Rostrum with 10-14 well developed spines dorsally, and 12-18 spines ventrally on the rostrum; ventral spines not restricted to distal half of rostrum...... 5

5. Carpus of second legs with 8-9 segments; median carapace teeth near rostral base fixed ...... Plesionika beebei Carpus of second legs with 14-17 segments; 3 median carapace teeth at rostral base moveable ...... Plesionika trispinus

6. Discoid widening of inner margin of ischium of first leg prominent...... Pandalopsis ampla No discoid widening of inner margin of ischium of first leg...... Pandalus 7

7. Carapace, abdomen and legs a uniform translucent pink; third abdominal somite with dorsal carina forming a lobe on posterior margin ...... Pandalus jordani Carapace or abdomen with white lines,spots or flecks, or if uniform pink, with yellow bands on legs; third abdominal somite not carinated dorsally...... 8

8. Carapace with white lines and abdomen with white spots OR legs with yellow bands; dorsal spines confined to anterior half of carapace...... 9 Carapace with scattered small black specks and minute red dots, abdomen with white lines dorsally and black lines laterally; dorsal spines extend to posterior half of carapace...... Pandalus danae

9. Carapace and abdomen uniform translucent pink, legs 3-5 pink with yellow bands; sixth abdominal somite length  2X width...... Pandalus tridens Carapace red with white lines, white spots on 1st and 5th abdominal somites, legs red and white banded; sixth abdominal somite length <2X width...... Pandalus platyceros REVISED KEY TO THE SO. CALIFORNIA BIGHT BENTHIC PENAEOID SHRIMP D. B. Cadien (LACSD)- 12 March 1998 (modified 27May98) (based on keys in Henrickx 1995, and Dall et al 1990)

1. Cervical sulcus reaching less than 2/3 the distance from the hepatic spine to the top of the carapace; postorbital spine absent; pereopod 4 lacks epipod...... 2 Cervical sulcus reaching the top of the carapace; postorbital spine present; pereopod 4 bearing an epipod ...... Solenoceridae 3

2. Third to 5th pleopods biramous; prosartema (eye brush) present; exopods on 2nd and 3rd maxillipeds ...... Penaeidae 4 Third to 5th pleopods uniramous; prosartema absent; exopods lacking on 2nd and 3rd maxillipeds ...... Sicyoniidae 6

3. Petasma of male with medial projection truncate and lateral projection acute and elongate (return to lab for identification) ...... Solenocera florea* Petasma of male with medial projection elongate and triangular, lateral projection short and blunt (return to lab for identification)...... Solenocera mutator

4. Rostrum dentate both dorsally and ventrally...... Penaeus 5 Rostrum dentate only dorsally...... Metapenaeopsis 6

5. Gastrofrontal carina well defined, reaching to orbital margin...... Penaeus californiensis Gastrofrontal carina often inconspicuous anteriorly, not reaching orbital margin...... Penaeus brevirostris*

6. Rostrum with 7-9 dorsal teeth, thelycum plate on sternite XIII of female lacking anteriorly directed spine ...... Metapenaeopsis kishinouyei*# Rostrum with 8-13 dorsal teeth, thelycum plate on sternite XIII of female bearing sharp anteriorly directed spine...... 7

7. Adult females with thelycum having a broad anteriomedially indented crest on sternite XIV (between 5th legs) ...... Metapenaeopsis mineri# Adult females with thelycum having a narrow anteriomedially cusped crest on sternite XIV (between 5th legs)...... Metapenaeopsis beebei*#

8. Carapace bearing lateral "bulls-eye" marking or brown spot inside a larger light area in the posterior mid-portion of the carapace ...... 9 Carapace without brown spot or other pronounced marking on the posterior mid-portion of the carapace......

9. Anterior dorsal tooth of first abdominal segment sub-equal in size to posterior-most tooth of carapace dorsal carina ...... Sicyonia penicillata Anterior dorsal tooth of first abdominal segment much larger than posterior-most tooth of carapace dorsal carina ...... Sicyonia disedwardsi*

*=not yet reported from area, but may range into it during strong ENSO events. Included for differentiation from closely related species known to occur in the southern California Bight. #=adult males separable only on details of petasma structure (consult Hendrickx 1995) REVISED KEY TO THE ALPHEIDAE OF CALIFORNIA D. B. CADIEN (CSDLAC) - 23MAR 1998 (based on the key in Wicksten 1984)

1. Triangular movable plate articulated at posterolateral angle of sixth abdominal somite lateral to uropod base...... 2 No triangular plate lateral to uropod base...... 9 2. Rostrum prominent, orbital hoods armed with spines ...... Alphaeopsis equidactylus Rostrum absent, carapace front without spines...... Betaeus 3 3. Dactyls of walking legs slender and simple...... 4 Dactyls of walking legs stout and bifid...... 6 4. Chelae of first legs with fingers > than palm; large male with fingers of chelipeds gaping ...... Betaeus longidactylus Chelae of first legs with fingers  than palm; large male with fingers of chelipeds not gaping ...... 5 5. Blade of antennal scale broad distally; fixed finger of first cheliped decreasing in width evenly to sharp curved tip...... Betaeus harrimani Blade of antennal scale narrow distally; fixed finger of first cheliped truncate before sharp curved tip ...... Betaeus ensenadensis 6. Carapace front curved, not emarginate; commensal with sea urchins (Strongylocentrotus spp) ...... Betaeus macginitieae Carapace front emarginate; commensal with abalone or free living ...... 7 7. Emargination of front shallow; telson with posterolateral spines small or missing; commensal with abalones (Haliotis spp)...... Betaeus harfordi Emargination of front deep; telson with posterolateral spines well developed...... 8 8. Peduncle of first antenna less than ½ carapace length; merus of cheliped with lower inner ridge with long bristles, upper ridge ending in sharp tooth; chela with fingers subequal to palm; chela 3 times as long as wide ...... Betaeus gracilis Peduncle of first antenna  carapace length; merus of cheliped with lower inner ridge usually tuberculate, upper ridge with tuft of hair; chela with fingers longer than palm; chela twice as long as wide...... Betaeus setosus 9. Eyes partially or fully exposed dorsally...... 10 Eyes fully covered by carapace dorsally ...... 12 10. Eyes fully exposed dorsally, rostrum shorter than eyestalks...... Automate 11 Eyes partially exposed dorsally, rostrum much longer than eyestalks ...... Salmoneus sp A 11. Propodus of 3rd legs bearing spines on the posterior margin ...... Automate dolichognatha Propodus of 3rd legs setose, but lacking spines on posterior margin...... Automate sp A 12. Legs lacking epipods; dactyls of legs 3-5 bifid...... Synalpheus lockingtoni Legs with epipods; dactyls of legs 3-5 simple ...... Alpheus 13 13. Dactyl of major chela closing horizontally; merus of 3rd leg with prominent inferior spine ...... Alpheus clamator Dactyl of major chela closing vertically; merus of 3rd leg lacking prominent inferior spine...... 14 14. Orbital hoods spined; minor chela with prominent spine posterior to movable finger; movable finger flattened (lamellate) ...... Alpheus bellimanus Orbital hoods not spined; minor chela without prominent spine posterior to movable finger; movable finger not flattened...... Alpheus californiensis